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''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
coelophysid Coelophysidae is a family of primitive carnivorous theropod dinosaurs. Most species were relatively small in size. The family flourished in the Late Triassic and Early Jurassic periods, and has been found on numerous continents. Many members of C ...
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
that lived approximately 228 to 201.3
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago) ...
during the latter part of the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
from the Carnian and Rhaetian faunal stages in what is now the southwestern United States. ''
Megapnosaurus ''Megapnosaurus'' (meaning "big dead lizard", from Greek μεγα = "big", 'απνοος = "not breathing", "dead", σαυρος = "lizard") is an extinct genus of coelophysid theropod dinosaur that lived approximately 188 million years ago dur ...
'' was once considered a species within this genus,Weishampel, David B; et al. (2004). "Dinosaur distribution (Early Jurassic, Africa)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka (eds.): The Dinosauria, 2nd, Berkeley: University of California Press. Pp. 535–536. but this interpretation has been challenged since 2017 and the genus ''Megapnosaurus'' is now considered valid. ''Coelophysis'' was a small, slenderly-built, ground-dwelling,
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
that could grow up to long. It is one of the earliest known dinosaur genera. Scattered material representing similar animals has been found worldwide in some
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
and
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
formations. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
''C. bauri'', originally given to the genus ''
Coelurus ''Coelurus'' ( ) is a genus of coelurosaurian dinosaur from the Late Jurassic period (mid-late Kimmeridgian faunal stage, 155–152 million years ago). The name means "hollow tail", referring to its hollow tail vertebrae ( Greek κο ...
'' by Edward Drinker Cope in 1887, was described by the latter in 1889. The names ''Longosaurus'' and ''Rioarribasaurus'' are synonymous with ''Coelophysis''. ''Coelophysis'' is one of the most specimen-rich dinosaur genera.


History of discovery

The type species of ''Coelophysis'' was originally named as a species of ''Coelurus''. Edward Drinker Cope first named ''Coelophysis'' in 1889 to name a new genus, outside of ''Coelurus'' and ''
Tanystropheus ''Tanystropheus'' (Greek ~ 'long' + 'hinged') is an extinct archosauromorph reptile from the Middle and Late Triassic epochs. It is recognisable by its extremely elongated neck, which measured long—longer than its body and tail combined. T ...
,'' which ''C. bauri'' was previously classified in, for ''C. bauri'', ''C. willistoni'' and ''C. longicollis''. An amateur
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
collector working for Cope, David Baldwin, had found the first remains of the dinosaur in 1881 in the Chinle Formation in northwestern New Mexico. Early in 1887, Cope referred the specimens collected to two new species, ''C. bauri'' and ''C. longicollis'' of the genus ''Coelurus''. Later in 1887 Cope reassigned the material to a yet another genus, ''Tanystropheus''. Two years later, Cope corrected his classification after realizing differences in the
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
e, and named ''Coelophysis'', with ''C. bauri'' as the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
, which was named for Georg Baur, a comparative anatomist whose ideas were similar to Cope's. The name ''Coelophysis'' comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words κοῖλος/koilos (meaning 'hollow') and φύσις/physis (meaning 'form'), together "hollow form", which is a reference to its hollow vertebrae. However, the first finds were too poorly preserved to give a complete picture of the new dinosaur. In 1947, a substantial 'graveyard' of ''Coelophysis'' fossils was found by George Whitaker, the assistant of Edwin H. Colbert, in
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
, at the
Ghost Ranch Ghost Ranch is a retreat and education center located close to the village of Abiquiú in Rio Arriba County in north central New Mexico, United States. It was the home and studio of Georgia O'Keeffe, as well as the subject of many of her painti ...
, close to the original find. American Museum of Natural History
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Edwin H. Colbert conducted a comprehensive study of all the fossils found up to that date and assigned them to ''Coelophysis''. The Ghost Ranch specimens were so numerous, including many well-preserved and fully articulated specimens, that one of them has since become the diagnostic, or
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
, for the entire genus, replacing the original, poorly preserved specimen.''"Syntarsus" rhodesiensis'' was first described by Raath (1969) and assigned to Podokesauridae.Raath (1969). "A new Coelurosaurian dinosaur from the Forest Sandstone of Rhodesia." Arnoldia Rhodesia. 4 (28): 1–25. The taxon "Podokesauridae", was abandoned because its type specimen was destroyed in a fire and can no longer be compared to new finds. Over the years paleontologists assigned the genus to
Ceratosauridae Ceratosauridae is an extinct family of theropod dinosaurs belonging to the infraorder Ceratosauria. The family's type genus, ''Ceratosaurus'', was first found in Jurassic rocks from North America. Ceratosauridae is made up of the genera ''Cer ...
(Welles, 1984), Procompsognathidae (Parrish and Carpenter, 1986) and
Ceratosauria Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with ''Ceratosaurus'' than with birds. The oldest known ceratosaur, '' Saltriovenator'', dates to the earliest ...
(Gauthier, 1986). In 2004 "Syntarsus" was found to be synonymous with ''Coelophysis'' by Tykoski and Rowe (2004). Ezcurra and Novas (2007) and Ezcurra (2007) also concluded that "Syntarsus" was synonymous with ''Coelophysis''.Tykoski, R. S., and Rowe, T., 2004, Ceratosauria, Chapter Three: In: The Dinosauria, Second Edition, edited by Weishampel, D.B., Dodson, P., and Osmolska, H., California University Press, p. 47-70. In a phylogenetic analysis by Ezcurra (2017), ''
Megapnosaurus ''Megapnosaurus'' (meaning "big dead lizard", from Greek μεγα = "big", 'απνοος = "not breathing", "dead", σαυρος = "lizard") is an extinct genus of coelophysid theropod dinosaur that lived approximately 188 million years ago dur ...
'' was recovered in a clade with '' Segisaurus'' and ''
Camposaurus ''Camposaurus'' ( ) is a coelophysid dinosaur genus from the Norian stage of the Late Triassic period of North America. The pertinent fossil remains date back to the early to middle Norian stage, and is widely regarded as the oldest known neot ...
'', supporting the generic distinction of ''Megapnosaurus''. This was supported by Barta and colleagues in 2018, noting that ''Coelophysis'' still bears the vestigial 5th metacarpal, a feature absent in ''Megapnosaurus''.


Description

''Coelophysis'' is known from a number of complete fossil skeletons of the species ''C. bauri'': a lightly built dinosaur that measured up to in length and was more than a meter tall at the hips. Paul (1988) estimated the weight of the gracile form at , and the weight of the robust form at , but later presented a higher estimate of . ''Coelophysis'' was a bipedal, carnivorous, theropod dinosaur and a fast and agile runner. Despite being an early dinosaur, the evolution of the
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
body form had already advanced greatly from creatures like ''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
'' and ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwes ...
''. The torso of ''Coelophysis'' conforms to the basic theropod body shape, but the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
displays some special characteristics: ''C. bauri'' had a
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
(wishbone), the earliest known example in a dinosaur. ''Coelophysis'' also preserves the ancestral condition of possessing four digits on the hand (manus). It had only three functional digits, the fourth embedded in the flesh of the hand. ''Coelophysis'' had narrow hips, forelimbs adapted for grasping prey, and narrow feet.Tykoski, R.S. & Rowe, T. (2004). "Ceratosauria": In: Its neck and tail were long and slender. The pelvis and hindlimbs of ''C. bauri'' are also slight variations on the theropod body plan. It has the open acetabulum and straight ankle hinge that define the Dinosauria. The hindlimb ended in a three-toed foot ( pes), with a raised
hallux Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being '' digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being '' pl ...
. The tail had an unusual structure within its interlocking
prezygapophysis The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...
of its vertebrae, which formed a semi-rigid lattice, apparently to stop the tail from moving up and down. ''Coelophysis'' had a long narrow head (approximately ), with large, forward-facing eyes that afforded it stereoscopic vision and as a result excellent depth perception. Rinehart ''et al.'' (2004) described the complete
sclerotic ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
found for a juvenile ''Coelophysis bauri'' (specimen NMMNH P-4200), and compared it to data on the sclerotic rings of reptiles and birds and concluded that ''Coelophysis'' was a diurnal, visually oriented predator. The study found that the vision of ''Coelophysis'' was superior to most lizards' vision, and ranked with that of modern birds of prey. The eyes of ''Coelophysis'' appear to be the closest to those of eagles and hawks, with a high power of accommodation. The data also suggested poor night vision, which would mean this dinosaur had a round rather than a split pupil. ''Coelophysis'' had an elongated snout with large fenestrae which helped to reduce skull weight, while narrow struts of bones preserved the structural integrity of the skull. The neck had a pronounced
sigmoid Sigmoid means resembling the lower-case Greek letter sigma (uppercase Σ, lowercase σ, lowercase in word-final position ς) or the Latin letter S. Specific uses include: * Sigmoid function, a mathematical function * Sigmoid colon, part of the l ...
curve. The braincase is known in ''Coelophysis bauri'' but little data could be derived because the skull was crushed. Unlike some other theropods, the cranial ornamentation of ''Coelophysis'' was not located at the top of its skull. Low, laterally raised bony ridges were present on the dorsolateral margin of the nasal and lacrimal bones in the skull, directly above the
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, bird ...
.


Distinguishing anatomical features

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism or group. According to Ezcurra (2007), and Bristowe and Raath (2004) ''Coelophysis'' can be distinguished based on the following features: the absence of an offset rostral process of the maxilla; the quadrate is strongly caudally; a small external mandibular fenestra, which is 9–10% of the mandibular length; and the anteroposterior length of the ventral lacrimal process is greater than 30% of its height. Several paleontologists consider ''Coelophysis bauri'' to be the same dinosaur as ''Megapnosaurus rhodesiensis'' (formerly ''Syntarsus''), however this has been refuted by the following: Downs (2000) concluded that ''C. bauri'' differs from ''C. rhodesiensis'' in cervical length, proximal and distal hindlimb proportions and proximal caudal vertebral anatomy;Downs, A. (2000). "''Coelophysis bauri'' and ''Syntarsus rhodesiensis'' compared, with comments on the preparation and preservation of fossils from the Ghost Ranch Coelophysis quarry": In: Tykoski and Rowe (2004) concluded that ''C. bauri'' differs from ''M. rhodesiensis'' in that it lacks a pit at the base of the nasal process of the premaxilla; Bristowe and Raath (2004) concluded that ''C. bauri'' differs from ''M. rhodesiensis'' in having a longer maxillary tooth row; Barta ''et al''. (2018) concluded that ''C. bauri'' differed from ''M. rhodesiensis'' in that it bears its 5th metacarpal; and several features in the musculature of the limbs according to Griffin (2018).Griffin, Christopher. (2018). Developmental patterns and variation among early theropods. Journal of Anatomy. 232. 604-640. 10.1111/joa.12775.


Classification

''Coelophysis'' is a distinct taxonomic unit (genus), composed of one species; '' C. bauri.'' Two additional originally described species, ''C. longicollis'' and ''C. willistoni'', are now considered dubious and undiagnostic. ''M. rhodesiensis'' was referred to ''Coelophysis'' for several years, but it is likely its own genus, and is known from the early Jurassic of southern Africa. A third possible species is ''Coelophysis kayentakatae'', previously referred to the genus ''Megapnosaurus,'' from the
Kayenta Formation The Kayenta Formation is a geological formation in the Glen Canyon Group that is spread across the Colorado Plateau province of the United States, including northern Arizona, northwest Colorado, Nevada, and Utah. Traditionally has been suggested ...
of the southwestern US. In recent phylogenetic analyses, ''"Syntarsus" kayentakatae'' has been shown to be distantly related to ''Coelophysis'' and ''Megapnosaurus'', suggesting that it belongs to its own genus. In the early 1990s, there was debate over the diagnostic characteristics of the first specimens collected, compared to the material excavated at the Ghost Ranch ''Coelophysis'' quarry. Some paleontologists were of the opinion that the original specimens were not diagnostic beyond themselves and, therefore, that the name ''C. bauri'' could not be applied to any additional specimens. They therefore applied a different name, ''Rioarribasaurus'', to the Ghost Ranch quarry specimens. Since the numerous well-preserved Ghost Ranch specimens were used as ''Coelophysis'' in most of the scientific literature, the use of ''Rioarribasaurus'' would have been very inconvenient for researchers, so a petition was given to have the type specimen of ''Coelophysis'' transferred from the poorly preserved original specimen to one of the well-preserved Ghost Ranch specimens. This would make ''Rioarribasaurus'' a definite synonym of ''Coelophysis'', specifically a junior objective synonym. In the end, the International Commission on Zoological Nomenclature (ICZN) voted to make one of the Ghost Ranch samples the actual type specimen for ''Coelophysis'' and dispose of the name ''Rioarribasaurus'' altogether (declaring it a ''nomen rejectum'', or "rejected name"), thus resolving the confusion. The name ''Coelophysis'' therefore became a ''nomen conservandum'' ("conserved name"). In a situation affecting many dinosaur taxa, some more recently discovered fossils were originally classified as new genera but may be species of ''Coelophysis''. For example, Prof. Mignon Talbot's 1911 discovery which she named ''Podokesaurus holyokensis'', has long been considered to be related to ''Coelophysis'', and some modern scientists consider ''Podokesaurus'' a synonym of ''Coelophysis''. Another specimen from the Portland Formation of the Hartford Basin, now at the
Boston Museum of Science The Museum of Science (MoS) is a science museum and indoor zoo in Boston, Massachusetts, located in Science Park, a plot of land spanning the Charles River. Along with over 700 interactive exhibits, the museum features a number of live presentat ...
, has also been referred to ''Coelophysis''. The specimen consists of sandstone casts of a pubis, tibia, three ribs, and a possible vertebra, and probably originated in a quarry in Middletown, Connecticut. However, both the type specimen of ''Podokesaurus'' and the Middletown specimen are typically considered indeterminate theropods today. Sullivan & Lucas (1999) referred one specimen from Cope's original material of ''Coelophysis'' (AMNH 2706) to what they thought was a newly discovered theropod, '' Eucoelophysis''. However, subsequent studies have shown that ''Eucoelophysis'' was misidentified, and is actually a primitive, non-dinosaurian
ornithodira Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinos ...
n closely related to '' Silesaurus''. The genus ''Syntarsus'' was named by Raath in 1969 for the type species '' Syntarsus rhodesiensis'' from Africa, and later applied to the North American '' Syntarsus kayentakatae''. It was renamed by American entomologist Dr. Michael Ivie ( Montana State University of Bozeman), Polish Australian Dr. Adam Ślipiński, and Polish Dr. Piotr Węgrzynowicz (Muzeum Ewolucji Instytutu Zoologii PAN of
Warsaw Warsaw ( pl, Warszawa, ), officially the Capital City of Warsaw,, abbreviation: ''m.st. Warszawa'' is the capital and largest city of Poland. The metropolis stands on the River Vistula in east-central Poland, and its population is officia ...
), the three scientists who discovered that the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
name ''Syntarsus'' was already taken by a colydiine
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
described in 1869. Many paleontologists did not like the naming of ''Megapnosaurus'', partially because taxonomists are generally expected to allow original authors of a name to correct any mistakes in their work. Raath was aware of the homonymy between the dinosaur ''Syntarsus'' and beetle ''Syntarsus'', but the group who published ''Megapnosaurus'' were led to believe Raath was deceased and therefore unable to correct his mistake, and proceeded accordingly. Mortimer (2012) pointed out that "Paleontologists might have reacted more positively if the replacement name (''Megapnosaurus'') hadn't been facetious, translating to "big dead lizard". Yates (2005) analyzed ''Coelophysis'' and ''Megapnosaurus'' and concluded that the two genera are almost identical, and suggested that ''Megapnosaurus'' was possibly synonymous with ''Coelophysis''. In 2004, Raath co-authored two papers in which he argued that ''Megapnosaurus'' (formerly ''Syntarsus'') was a junior synonym of ''Coelophysis''. ''
Megapnosaurus ''Megapnosaurus'' (meaning "big dead lizard", from Greek μεγα = "big", 'απνοος = "not breathing", "dead", σαυρος = "lizard") is an extinct genus of coelophysid theropod dinosaur that lived approximately 188 million years ago dur ...
'' was regarded by Paul (1988) and Downs (2000) as being
congeneric Congener may refer to: * A thing or person of the same kind as another, or of the same group. * Congener (biology), organisms within the same genus. * Congener (chemistry), related chemicals, e.g., elements in the same group of the periodic table. ...
with ''Coelophysis''. Then in 1993, Paul suggested that ''Coelophysis'' should be placed in ''Megapnosaurus'' (then known as ''Syntarsus'') to get around the above-mentioned taxonomic confusion.Paul G.S. (1993). "Are ''Syntarsus'' and the Whitetaker theropod the same genus?": In: Downs (2000) examined ''Camposaurus'' and concluded that it was a junior synonym of ''Coelophysis'', because of its similarity to some of the ''Coelophysis'' Ghost Ranch specimens. However, a reassessment of the ''Camposaurus'' holotype by Martin Ezcurra and Stephen Brusatte published in 2011 revealed a pair of
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
in the holotype, indicating that ''C. arizonensis'' was not a synonym of ''C. bauri'', although it was a close relative of ''M. rhodesensis''. Barta ''et al''. (2018) concluded that ''C. bauri'' differed from ''M. rhodesiensis'' in that it bears its 5th metacarpal and several features in the musculature of the limbs according to Griffin (2018).


Paleobiology


Feeding

The teeth of ''Coelophysis'' were typical of predatory dinosaurs, blade-like, recurved, sharp and jagged with fine serrations on both the anterior and posterior edges. Its dentition shows that it was carnivorous, probably preying on the small, lizard-like animals that were discovered with it. It may also have hunted in packs to tackle larger prey. ''Coelophysis bauri'' has approximately 26 teeth on the maxillary bone of the upper jaw and 27 teeth on the dentary bone of the lower jaw. Carpenter (2002) examined the bio-mechanics of theropod forelimbs and attempted to evaluate their usefulness in predation. He concluded that the forelimb of ''Coelophysis'' was flexible and had a good range of motion, but its bone structure suggested that it was comparatively weak. The "weak" forelimbs and small teeth in this genus, suggested that ''Coelophysis'' preyed upon animals that were substantially smaller than itself. Rinehart ''et al.'' agreed that ''Coelophysis'' was a "hunter of small, fast-moving prey". Carpenter also identified three distinct models of theropod forelimb use and noted that ''Coelophysis'' was a "combination grasper-clutcher" as compared to other dinosaurs that were "clutchers" or "long armed graspers". It has been suggested that ''C. bauri'' was a
cannibal Cannibalism is the act of consuming another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded in more than 1,500 species. Human cannibalism is well documented, bo ...
, based on supposed juvenile specimens found "within" the abdominal cavities of some Ghost Ranch specimens. However, Robert J. Gay showed in 2002 that these specimens were misinterpreted. Several specimens of "juvenile coelophysids" were actually small crurotarsan reptiles such as '' Hesperosuchus''. Gay's position was lent support in a 2006 study by Nesbitt ''et al.'' In 2009, new evidence of cannibalism came to light when additional preparation of previously excavated matrix revealed regurgitate material in and around the mouth of ''Coelophysis'' specimen NMMNH P-44551. This material included tooth and jaw bone fragments that Rinehart ''et al.'' considered "morphologically identical" to a juvenile ''Coelophysis.'' In 2010, Gay examined the bones of juveniles found within the thoracic cavity of AMNH 7224, and calculated that the total volume of these bones was 17 times greater than the maximum estimated stomach volume of the ''Coelophysis'' specimen. Gay observed that the total volume would be even greater when considering that there would have been flesh on these bones. This analysis also noted the absence of tooth marks on the bones as would be expected in defleshing, and the absence of expected pitting by stomach acids. Finally, Gay demonstrated that the alleged cannibalized juvenile bones were deposited stratigraphically below the larger animal that had supposedly cannibalized them. Taken together these data suggested that the ''Coelophysis'' specimen AMNH 7224 was not a cannibal and that the bones of the juvenile and adult specimens were found in their final position as a result of "coincidental superposition of different sized individuals.


Pack behavior

The discovery of over 1000 specimens of ''Coelophysis'' at the Whitaker quarry at
Ghost Ranch Ghost Ranch is a retreat and education center located close to the village of Abiquiú in Rio Arriba County in north central New Mexico, United States. It was the home and studio of Georgia O'Keeffe, as well as the subject of many of her painti ...
, has suggested gregarious behavior to researchers like Schwartz and Gillette. There is a tendency to see this massive congregation of animals as evidence for huge packs of ''Coelophysis'' roaming the land. No direct evidence for flocking exists; the deposits only indicate that large numbers of ''Coelophysis'', along with other Triassic animals, were buried together. Some of the evidence from the
taphonomy Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
of the site indicates that these animals may have been gathered together to feed or drink from a depleted water hole or to feed on a
spawning Spawn is the eggs and sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of releasing the eggs and sperm, and the act of both sexes is called spawning. Most aquatic animals, except for aquat ...
run of fish, and then became buried in a catastrophic flash flood or a drought. With 30 specimens of ''C. rhodesiensis'' found together in Zimbabwe some palaeontologists have suggested that ''Coelophysis'' was indeed gregarious. Again there is no direct evidence of flocking in this case and it has also been suggested that these individuals were also victims of flash flooding as it appears to have been commonplace during this period.G. Bond, 1965. Some new fossil localities in the Karroo System of Rhodesia. Arnoldia, Series of Miscellaneous Publications, National Museum of Southern Rhodesia 2(11):1–4M. A. Raath, 1977. The Anatomy of the Triassic Theropod Syntarsus rhodesiensis (Saurischia: Podokesauridae) and a Consideration of Its Biology. Department of Zoology and Entomology, Rhodes University, Salisbury, Rhodesia 1–233


Growth and sexual dimorphism

Rinehart (2009) assessed the ontogenic growth of this genus using data gathered from the length of its upper leg bone (
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
) and concluded that ''Coelophysis'' juveniles grew rapidly, especially during the first year of life. ''Coelophysis'' likely reached sexual maturity between the second and third year of life and reached its full size, just above 10 feet in length, by its eighth year. This study identified four distinct growth stages: 1-year, 2-year, 4-year, and 7+ year. It was also thought, that as soon as they were hatched, they would have to fend for themselves. Two "morphs" of ''Coelophysis'' have been identified: a more gracile form, as in specimen AMNH 7223, and a slightly more
robust Robustness is the property of being strong and healthy in constitution. When it is transposed into a system, it refers to the ability of tolerating perturbations that might affect the system’s functional body. In the same line ''robustness'' ca ...
form, as in specimens AMNH 7224 and NMMNH P-42200. Skeletal proportions were different between these two forms; the gracile form has a longer skull, a longer neck, shorter forelimbs, and has
sacral Sacral may refer to: *Sacred Sacred describes something that is dedicated or set apart for the service or worship of a deity; is considered worthy of spiritual respect or devotion; or inspires awe or reverence among believers. The property ...
neural spines that are fused; and the robust form has a shorter skull, a shorter neck, longer forelimbs, and unfused sacral neural spines. Historically, many arguments have been made that this represents some sort of dimorphism in the population of ''Coelophysis'', probably
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
.Gay, R. (2005). "Sexual Dimorphism in the Early Jurassic Theropod Dinosaur ''
Dilophosaurus ''Dilophosaurus'' ( ) is a genus of theropod dinosaurs that lived in what is now North America during the Early Jurassic, about 193 million years ago. Three skeletons were discovered in northern Arizona in 1940, and the two best preserve ...
'' and a Comparison with Other Related Forms": In:
Raath agreed that dimorphism in ''Coelophysis'' is evidenced by the size and structure of the forelimb. Rinehart ''et al.'' studied 15 individuals, and agreed that two morphs were present, even in juvenile specimens, and suggested that sexual dimorphism was present early in life, prior to sexual maturity. Rinehart concluded that the gracile form was female and the robust form was male based on differences in the sacral vertebrae of the gracile form, which allowed for greater flexibility for egg laying. Further support for this position was provided by an analysis showing that each morph comprised 50% of the population, as would be expected in a 50/50 sex ratio. However, more recent research has found that ''C. bauri'' and ''C. rhodesiensis'' had highly variable growth between individuals, with some specimens being larger in their immature phase than smaller adults were when completely mature; this indicates that the supposed presence of distinct morphs is simply the result of individual variation. This highly variable growth was likely ancestral to dinosaurs but later lost, and may have given such early dinosaurs an evolutionary advantage in surviving harsh environmental challenges.


Reproduction

Through the compilation and analysis of a database of nearly three dozen birds and reptiles, and comparison with existing data about the anatomy of ''Coelophysis'' Rinehart ''et al.'' (2009) drew the following conclusions. It was estimated that average egg of ''Coelophysis'' was 31–33.5 millimeters across its minor diameter, and that each female would lay between 24 and 26 eggs in each clutch. The evidence suggested that some parental care was necessary to nurture the relatively small hatchlings during the first year of life, where they would reach 1.5 meters in length by the end of their first growth stage. ''Coelophysis bauri'' invested as much energy in reproduction as other extinct reptiles of its approximate size.


Paleopathology

In a 2001 study conducted by Bruce Rothschild and other paleontologists, 14-foot bones referred to ''Coelophysis'' were examined for signs of
stress fracture A stress fracture is a fatigue-induced bone fracture caused by repeated stress over time. Instead of resulting from a single severe impact, stress fractures are the result of accumulated injury from repeated submaximal loading, such as running or ...
, but none were found.Rothschild, B.; Tanke, D.H. & Ford, T.L. (2001). "Theropod stress fractures and tendon avulsions as a clue to activity": In: In ''C. rhodesiensis'', healed fractures of the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
and
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
have been observed, but are very rare. " e supporting butresses of the second sacral rib" in one specimen of ''Syntarsus rhodesiensis'' showed signs of fluctuating asymmetry. Fluctuating asymmetry results from developmental disturbances and is more common in populations under stress and can therefore be informative about the quality of conditions a dinosaur lived under.Molnar, R. E., 2001, Theropod paleopathology: a literature survey: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 337-363.


Ichnology

Edwin H. Colbert has suggested that the theropod footprints referred to the
ichnogenus An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' and ταξις, ''taxis'' meaning ...
''
Grallator ''Grallator'' GRA-luh-tor"is an ichnogenus (form taxon based on footprints) which covers a common type of small, three-toed print made by a variety of bipedal theropod dinosaurs. ''Grallator''-type footprints have been found in formations dati ...
'', located in the
Connecticut River Valley The Connecticut River is the longest river in the New England region of the United States, flowing roughly southward for through four states. It rises 300 yards (270 m) south of the U.S. border with Quebec, Canada, and discharges at Long Island ...
across Connecticut and Massachusetts, may have been made by ''Coelophysis''. The footprints are from the Late Triassic to Early Jurassic aged Newark Supergroup. They clearly show digits II III and IV but not I or V. That condition is strange for footprints of their age. The digits I and V were presumed to be stubby and ineffective, not touching the ground when the dinosaur was walking or running. They have been thought to be from an unidentified, primitive saurischian similar to ''Coelophysis'' by David B. Weishampel and L. Young more recently. Skeletal remains resembling ''Coelophysis'' have also been found in the valley, supporting the idea that a species similar to ''Coelophysis'' is responsible for the footprints.


Paleoenvironment

Specimens of ''Coelophysis'' have been recovered from the Chinle Formation of New Mexico and Arizona, more famously at the Ghost Ranch (Whitaker) quarry in the Rock Point member among other quarries in the underlying Petrified Forest member, the sediments of which have been dated to approximately 212 million years ago, making them part of the middle Norian stage of the Late Triassic, but Thomas Holtz Jr. interpreted that it was during the Rhaetian stage from approximately 204 to 201.6 million years ago.Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2011 Appendix.
/ref> ''C. rhodesiensis'' has been recovered in the Upper Elliott Formation in the
Cape A cape is a clothing accessory or a sleeveless outer garment which drapes the wearer's back, arms, and chest, and connects at the neck. History Capes were common in medieval Europe, especially when combined with a hood in the chaperon. Th ...
and Free State provinces of South Africa as well as the Chitake River bonebed quarry at the Forest Sandstone Formation in Zimbabwe. Ghost Ranch was located close to the equator over 200 million years ago, and had a warm, monsoon-like climate with heavy seasonal precipitation. Hayden Quarry, a new excavation site at Ghost Ranch, New Mexico, has yielded a diverse collection of fossil material that included the first evidence of dinosaurs and less-advanced dinosauromorphs from the same time period. The discovery indicates that the two groups lived together during the early Triassic period 235 million years ago. Therrien and Fastovsky (2001) examined the paleoenvironment of ''Coelophysis'' and other early theropods from Petrified Forest National Park in Arizona, and determined that this genus lived in an environment that consisted of floodplains marked by distinct dry and wet seasons. There was a great deal of competition during drier times when animals struggled for water in riverbeds that were drying up. In the upper sections of the Chinle Formation where ''Coelophysis'' is found, dinosaurs were rare; so far only ''
Chindesaurus ''Chindesaurus'' ( ) is an extinct genus of basal saurischian dinosaur from the Late Triassic (213-210 million years ago) of the southwestern United States. It is known from a single species, ''C. bryansmalli'', based on a partial skeleton recovere ...
'' and ''
Daemonosaurus ''Daemonosaurus'' is an extinct genus of possible theropod dinosaur from the Late Triassic of New Mexico. The only known fossil is a skull and neck fragments from deposits of the latest Triassic Chinle Formation at Ghost Ranch. ''Daemonosaurus'' ...
'' are known, the terrestrial fauna being dominated instead by other reptiles like the rhynchocephalian ''
Whitakersaurus ''Whitakersaurus'' is a genus of sphenodontid rhynchocephalian reptile dated to be late Triassic in age and is from the Ghost Ranch fossil quarry in New Mexico, USA. It is named after the discoverer of the Ghost Ranch quarry, George O. Whitaker ...
''; the
pseudosuchian Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
''
Revueltosaurus ''Revueltosaurus'' ("Revuelto lizard") is an extinct genus of suchian pseudosuchian from Late Triassic (late Carnian to middle Norian stage) deposits of New Mexico, Arizona and North Carolina, United States. Many specimens, mostly teeth, have bee ...
''; the
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order (biology), order Aetosauria (; from Ancient Greek, Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized Omnivore, omnivorous or Herbivore, herbivoro ...
s ''
Desmatosuchus ''Desmatosuchus'' (, from Greek δεσμός ''desmos'' 'link' + σοῦχος ''soûkhos'' 'crocodile') is an extinct genus of archosaur belonging to the Order Aetosauria. It lived during the Late Triassic. Description ''Desmatosuchus'' was ...
'', '' Typothorax'' and '' Heliocanthus''; the crocodilomorph '' Hesperosuchus''; the "
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
ns" ''
Shuvosaurus ''Shuvosaurus'' (meaning "Shuvo's lizard") is a genus of beaked reptile from the Late Triassic of western Texas. Despite looking superficially similar to a theropod dinosaur, it is actually more closely related to crocodilians. Discovery and ...
'',Irmis, R.B. (2005). "The vertebrate fauna of the Upper Triassic Chinle Formation in northern Arizona": In: ''
Effigia ''Effigia'' was an Extinction, extinct genus of Shuvosauridae, shuvosaurid known from the Late Triassic of New Mexico, south-western USA. With a bipedal stance, long neck, and a toothless beaked skull, ''Effigia'' and other shuvosaurids bore a re ...
'' and '' Vivaron''; along other rare components like the dinosauriform '' Eucoelophysis'', and the amniote '' Kraterokheirodon''. In the waterways there are the
phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent g ...
'' Machaeroprosopus''; the
archosauromorph Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, l ...
'' Vancleavea;'' the amphibians '' Apachesaurus'' and '' Koskinonodon''; and the fishes '' Reticulodus'', '' Arganodus'' and '' Lasalichthyes''.


Taphonomy

The multitude of specimens deposited so closely together at Ghost Ranch was probably the result of a flash flood, which swept away a large number of ''Coelophysis'' and buried them quickly and simultaneously. In fact, it seems that such flooding was commonplace during this period of the
Earth's history The history of Earth concerns the development of planet Earth from its formation to the present day. Nearly all branches of natural science have contributed to understanding of the main events of Earth's past, characterized by constant geologic ...
and, indeed, the
Petrified Forest Petrified wood, also known as petrified tree (from Ancient Greek meaning 'rock' or 'stone'; literally 'wood turned into stone'), is the name given to a special type of '' fossilized wood'', the fossilized remains of terrestrial vegetation. ' ...
of nearby
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
is the result of a preserved log jam of tree trunks that were caught in one such flood. Whitaker quarry at Ghost Ranch is considered a monotaxic site because it features multiple individuals of a single taxon. The quality of preservation and the ontogenic (age) range of the specimens helped make ''Coelophysis'' one of the best known of all genera. In 2009, Rinehart ''et al.'' noted that in one case the ''Coelophysis'' specimens were "washed into a topographic low containing a small pond, where they probably drowned and were buried by a sheet flood event from a nearby river." The 30 specimens of ''C. rhodesiensis'' found together in Zimbabwe was also probably the result of a flash flood, which swept away a large number of ''Coelophysis'' and buried them quickly and simultaneously as well.


Cultural significance

''Coelophysis'' was the second dinosaur in space, following ''
Maiasaura ''Maiasaura'' (from the Greek ''μαῖα'', meaning "good mother" and ''σαύρα'', the feminine form of ''saurus'', meaning "reptile") is a large herbivorous saurolophine hadrosaurid ("duck-billed") dinosaur genus that lived in the area cur ...
'' (
STS-51-F STS-51-F (also known as Spacelab 2) was the 19th flight of NASA's Space Shuttle program and the eighth flight of Space Shuttle ''Challenger''. It launched from Kennedy Space Center, Florida, on July 29, 1985, and landed eight days later on Aug ...
). A ''Coelophysis'' skull from the
Carnegie Museum of Natural History The Carnegie Museum of Natural History (abbreviated as CMNH) is a natural history museum in the Oakland neighborhood of Pittsburgh, Pennsylvania. It was founded by Pittsburgh-based industrialist Andrew Carnegie in 1896. Housing some 22 million ...
was aboard the
Space Shuttle Endeavour Space Shuttle ''Endeavour'' ( Orbiter Vehicle Designation: OV-105) is a retired orbiter from NASA's Space Shuttle program and the fifth and final operational Shuttle built. It embarked on its first mission, STS-49, in May 1992 and its 25th and ...
mission
STS-89 STS-89 was a Space Shuttle mission to the '' Mir'' space station flown by Space Shuttle '' Endeavour'', and launched from Kennedy Space Center, Florida on 22 January 1998. Crew Crew notes STS-89 was originally scheduled to return Wendy B. La ...
when it left the atmosphere on 22 January 1998. It was also taken onto the space station
Mir ''Mir'' (russian: Мир, ; ) was a space station that operated in low Earth orbit from 1986 to 2001, operated by the Soviet Union and later by Russia. ''Mir'' was the first modular space station and was assembled in orbit from 1986 to&n ...
before being returned to Earth. As being over 100 years old, ''Coelophysis'' is one of the best-known dinosaurs in literature. It was designated as the official state fossil of New Mexico in 1981, and is now the logo of the New Mexico Museum of Natural History.


References


External links


''Coelophysis''
in the Dino Directory {{Portal bar, Paleontology, Dinosaurs Coelophysoids Late Triassic dinosaurs of North America Chinle fauna Triassic geology of New Mexico Paleontology in New Mexico Symbols of New Mexico Fossil taxa described in 1889 Taxa named by Edward Drinker Cope